REPORTS

However, the absence of abundant lithic assemblages at these Hata archaeology sites
requires explanation.
At the nearby Gona site, abundant Oldowan
tools were made and discarded immediately
adjacent to cobble conglomerates that offered
excellent, easily accessible raw materials for
stone-tool manufacture. It has been suggested
that the surprisingly advanced character of this
earliest Oldowan technology was conditioned
by the ease of access to appropriate fine-grained
raw materials at Gona (10). Along the Karari
escarpment at Koobi Fora (13), the basin margin at Fejej (14), and the lake margin at Olduvai
Gorge (12), hominids also had easy access to
nearby outcrops of raw material. In contrast, the
diminutive nature of the Oldowan assemblages
in the lower Omo [made on tiny quartz pebbles
(15)] was apparently conditioned by a lack of
available large clasts.
The situation on the Hata lake margin was
even more difficult for early toolmakers. Here,
raw materials were not readily available because
of the absence of streams capable of carrying
even pebbles. There were no nearby basalt outcrops. The absence of locally available raw material on the flat featureless Hata lake margin
may explain the absence of lithic artifact concentrations. The bone modification evidence
demonstrates that early hominids were transporting stone to the site of carcass manipulation.
The paucity of evidence for lithic artifact abandonment at these sites suggests that these early
hominids may have been curating their tools
(cores and flakes) with foresight for subsequent
use. Indications of tool curation by later hominids have been found at the more recent Pleistocene sites of Koobi Fora [Karari escarpment
versus Ileret (13)] and Swartkrans [polished
bone tools in a single repository (16)].
Additional research into the Hata beds
may allow a determination of whether the
butchery is related to hunting or scavenging.
The Bouri discoveries show that the earliest
Pliocene archaeological assemblages and
their landscape patterning are strongly conditioned by the availability of raw material.
They demonstrate that a major function of the
earliest known tools was meat and marrow
processing of large carcasses. Finally, they
extend this pattern of butchery by hominids
well into the Pliocene.
References and Notes

1. B. Asfaw et al., Science 284, xxx (1999).

2. N. J. Hayward and C. J. Ebinger, Tectonics 15, 244
(1996).
3. J. E. Kalb et al., Nature 298, 17 (1982). Eleven years
later, J. E. Kalb et al. [Newsl. Stratigr. 29, 21 (1993)]
followed Clark et al. (4) in reversing Daka/Bodo
Member order.
4. J. D. Clark et al., Nature 307, 423 (1984).
5. J. D. Clark et al., Science 264, 1907 (1994).
6. P. R. Renne, G. WoldeGabriel, W. K. Hart, G. Heiken,
T. D. White, Geol. Soc. Am. Bull., in press.
7. The age of this standard is now known to be slightly
older [28.02 Ma (17)], but comparison with previous
data (for example, from the Gona) is facilitated by

8.
9.

10.
11.

12.
13.
14.
15.
16.
17.
18.

retaining the standard age of 27.84 Ma. A table of the

Ar isotopic data is available at www.sciencemag.org/
feature/data/991110.shl.
F. J. Hilgen, Earth Planet. Sci. Lett. 107, 249 (1991).
The sedimentation rate inferred is conservative because comparison of the 40Ar/39Ar age with the age
of 2.6 Ma (8) for the Gauss/Matuyama boundary
more properly requires the use of the older age of the
standard (28.02 Ma) and indicates an ;10% greater
sedimentation rate.
S. Semaw et al., Nature 385, 333 (1997).
T. D. White, Prehistoric Cannibalism at Mancos
5MTUMR-2346 (Princeton Univ. Press, Princeton, NJ,
1992); L. R. Binford, Bones: Ancient Men and Modern
Myths (Academic Press, New York, 1981); R. J. Blumenschine, J. Hum. Evol. 29, 21 (1995); S. D. Capaldo
and R. J. Blumenschine, Am. Antiq. 59, 724 (1994).
R. J. Blumenschine and F. T. Masao, J. Hum. Evol. 21,
451 (1991).
G. L. Isaac, Ed., Koobi Fora Research Project Volume 5:
Plio-Pleistocene Archaeology (Clarendon, Oxford,
1997).
B. Asfaw et al., J. Hum. Evol. 21, 137 (1991).
F. C. Howell, P. Haeserts, J. de Heinzelin, ibid. 16, 665
(1987).
C. K. Brain, Ed., Swartkrans: A Caves Chronicle of Early
Man, vol. 8 of Transvaal Museum Monograph Series
(Transvaal Museum, Pretoria, 1993).
P. R. Renne et al., Chem. Geol. 145, 117 (1998).
The Middle Awash paleoanthropological project is
multinational (13 countries), with interdisciplinary
research codirected by B. Asfaw, Y. Beyene, J. D. Clark,
T. D. White, and G. WoldeGabriel. The research re-

ported here was supported by NSF, the Ann and

Gordon Getty Foundation (Berkeley Geochronology
Center), and the Institute of Geophysics and Planetary Physics of the University of California and the
Earth Environmental Sciences Division at Los Alamos
National Laboratory. Additional contributions were
made by the Graduate School, the Office for Advancement of Scholarship and Teaching, and the
Department of Geology at Miami University. We
thank the Ethiopian Mapping Agency and the NASA
Goddard Space Flight Center for imagery. We thank
H. Gilbert for fieldwork and for work on the illustrations. D. Brill made the photographs and G. Richards
and B. Plowman at the University of Pacific made the
scanning electron microscope (SEM) image. T. Larson
provided invaluable field and laboratory geology support. A. Defleur assisted in field survey and excavations at BOU-VP-11. We thank H. Saegusa for proboscidean identifications, D. DeGusta for primate
identifications and excavations at BOU-VP-12/1, and
F. C. Howell for carnivore identifications. We thank
O. Lovejoy, G. Suwa, and B. Asfaw for helpful comments. We thank the Ethiopian Ministry of Information and Culture, the Centre for Research and Conservation of the Cultural Heritage, and the National
Museum of Ethiopia. We thank the Afar Regional
Government and the Afar people of the Middle
Awash for permission and support. We thank the
many individuals who contributed to the camp,
transport, survey, excavation, and laboratory work
that stands behind the results presented.
25 February 1999; accepted 30 March 1999

Australopithecus garhi: A New

Species of Early Hominid
from Ethiopia
Berhane Asfaw,1 Tim White,2* Owen Lovejoy,3
Bruce Latimer,4,5 Scott Simpson,5 Gen Suwa6
The lack of an adequate hominid fossil record in eastern Africa between 2 and
3 million years ago (Ma) has hampered investigations of early hominid phylogeny. Discovery of 2.5 Ma hominid cranial and dental remains from the Hata
beds of Ethiopias Middle Awash allows recognition of a new species of Australopithecus. This species is descended from Australopithecus afarensis and is
a candidate ancestor for early Homo. Contemporary postcranial remains feature
a derived humanlike humeral/femoral ratio and an apelike upper armtolower
arm ratio.
The succession of early hominid genera and
species indicates diversification into at least
two distinct adaptive patterns by ;2.7 Ma. A
meager east African hominid record between
2 and 3 Ma has caused the pattern and pro1
Rift Valley Research Service, Post Office Box 5717,
Addis Ababa, Ethiopia. 2Laboratory for Human Evolutionary Studies, Museum of Vertebrate Zoology, and
Department of Integrative Biology, University of California at Berkeley, Berkeley, CA 94720, USA. 3Department of Anthropology and Division of Biomedical
Sciences, Kent State University, Kent OH 44242, USA.
4
Cleveland Museum of Natural History and 5Department of Anatomy, School of Medicine, Case Western
Reserve University, Cleveland, OH 44106, USA. 6University Museum, University of Tokyo, Hongo, Bunkyoku, Tokyo, 113-0033, Japan.

*To whom correspondence should be addressed. Email: timwhite@socrates.berkeley.edu

cess of this diversification to remain obscure.

The Australopithecus afarensis (3.6 to 3.0
Ma) to A. aethiopicus (2.6 Ma) to A. boisei
(2.3 to 1.2 Ma) species lineage is well corroborated by craniodental remains. In contrast, a suggested relationship between A. afarensis and early Homo has previously been
evidenced only by relatively uninformative
isolated teeth (1), a palate (2), and a temporal
fragment (3).
The recovery of hominid remains from the
Hata (abbreviation of Hatayae) Member of the
Bouri Formation adds substantially to the inventory of fossils bearing on these phylogenetic
issues. These remains comprise craniodental
and postcranial elements from several areas in
the Middle Awash. The first of these was discovered in 1990 at Matabaietu and Gamedah.

www.sciencemag.org SCIENCE VOL 284 23 APRIL 1999

629

REPORTS
Biochronology and Ar/Ar dating place these
remains at ;2.5 Ma. They include a small left
parietal fragment (GAM-VP-1/2) and an edentulous left mandible corpus from Gamedah
(GAM-VP-1/1), as well as a distal left humerus
from Matabaietu (MAT-VP-1/1). It is impossible to attribute the humerus and parietal fragments to a genus. However, the small, fluvially
abraded Gamedah mandible retains tooth roots
and corpus contours. These demonstrate that it
is not a robust Australopithecus.
It was not until 1996 1998 that we recovered additional hominid remains of comparable antiquity west of the modern Awash, at
Bouri. The proximal half of an adult hominid
ulna (BOU-VP-11/1) was found on the surface of the Hata beds by T. Assebework on 17
November 1996. On 30 November, White
found a proximal femur and associated forearm elements of a smaller individual, ;100
m to the WNW (BOU-VP-12/1A-G). Sieving
and excavation revealed additional portions
of this individuals femur in situ, 1 m above
a 2.496 Ma volcanic ash, in a horizon with
abundant catfish remains and medium-sized
bovid fossils, the latter bearing cut marks (4).
This partial hominid skeleton includes fairly
complete shafts of a left femur and the right
humerus, radius, and ulna. A partial fibular
shaft, a proximal foot phalanx, and the base of
the anterior portion of the mandible were also
found. There is no evidence that these remains
represent more than one individual. Except for
the in situ distal femoral shaft segment, all were
surface finds lying within 2 m of one another.
All are similarly preserved. Lengths can be
accurately estimated for the phalanx, the femur,
and the three arm elements. The foot phalanx is
similar to remains of A. afarensis in size,
length, and curvature. The mandible does not
retain diagnostic morphology. No associated
hominid teeth were found on the surface or in a
large excavation.
Further search of the same ;2.5 Ma horizon
led to the discovery, 278 m farther NNW, of a
partial hominid cranium (BOU-VP-12/130) on
20 November 1997 by Y. Haile-Selassie (Fig.
1). Another individuals crested cranial vault
fragment (BOU-VP-12/87) was found 50 m
south of the BOU-VP-12/1 skeleton excavation. At a more northerly locality in the Esa
Dibo area ;9 km away, A. Defleur found a
fairly complete mandible, with dentition, of
another hominid individual (BOU-VP-17/1) on
17 November 1997. An additional hominid humeral shaft (BOU-VP-35/1) was found ;1 km
farther north of the location of the mandible, on
4 December 1998, by D. DeGusta. On biochronological grounds, these Esa Dibo specimens
are about the same age as the more southerly
cluster of hominid remains at Bouri localities
11 and 12 (4).
Great uncertainty has continued to confound the origin of Homo because of a lack of
evidence from the interval between 2 and 3

630

Ma (5). The 2.5 Ma Bouri Hata hominids

bear directly on these issues. In addition, they
are closely associated with behavioral evidence of lithic technology (4). Australopithecus africanus from South Africa is roughly
contemporary with the Hata remains. In eastern Africa, A. aethiopicus and at least one
other putative lineage ancestral to early
Homo are contemporaries in the Turkana Basin. The BOU-VP-12/130 cranial remains
represent no previously named species. Only
the recovery of additional specimens with
associated crania and dentitions may allow
the Bouri postcrania to be positively attributed to this new taxon. Therefore, the new
species described below is established strictly
on the basis of craniodental remains.
The following is a description of Austra-

lopithecus garhi, based on the BOU-VP-12/

130 specimen: order, Primates Linnaeus
1758; suborder, Anthropoidea Mivart 1864;
genus, Australopithecus DART 1925; and
species, Australopithecus garhi.
Etymology. The word garhi means surprise in the Afar language.
Holotype. ARA-VP-12/130 is an associated
set of cranial fragments comprising the frontal,
parietals, and maxilla with dentition. It was
found by Y. Haile-Selassie on 20 November
1997. The holotype is housed at the National
Museum of Ethiopia, Addis Ababa.
Locality. Bouri Vertebrate Paleontology locality 12 (BOU-VP-12) is on the eastern side of the Bouri peninsula, west of the
modern Awash River, in the Middle Awash
paleoanthropological study area, Afar depres-

cm

Fig. 1. Cranial parts of BOU-VP-12/130. (Top) Superior view of the original fossil. Nonstandard
orientation (rotated posteriorly ;10 from Frankfurt horizontal) to show maximum anatomy. (Bottom)
Lateral view of casts to show cranial and maxillary profiles. Note that neither Frankfurt horizontal nor
placement of the maxilla relative to the vault can be accurately determined and that reconstructed
portions (indicated by oblique lines) are speculative. Photos David L. Brill 1999\Atlanta.

23 APRIL 1999 VOL 284 SCIENCE www.sciencemag.org

REPORTS
sion, Ethiopia. The BOU-VP-12/130 holotype
was found at 1015.61999N, 4033.84459E, at
;550 m elevation.
Horizon and associations. The holotype
was recovered from silty clays within 2 m of
the top of the Maoleem vitric tuff, which has
been dated to 2.496 Ma by Ar/Ar. Vertebrate
fossils, including additional hominids, were
found at the same stratigraphic horizon on
nearby outcrops (4).

Diagnosis. Australopithecus garhi is a

species of Australopithecus distinguished
from other hominid species by a combination
of characters presented in Table 1. It is distinguished from A. afarensis by its absolutely
larger postcanine dentition and an upper third
premolar morphology with reduced mesiobuccal enamel line projection and less occlusal asymmetry. Australopithecus garhi lacks
the suite of derived dental, facial, and cranial

Table 1. List of characters. Listed are characters widely used in consideration of

hominid phylogenetics (11, 13) that are preserved on the BOU-VP-12/130
holotype cranium. Because of arbitrary boundaries of presence or absence
criteria, variability within species, limited sample sizes, and possible correlation
between features, we caution against a numerical cladistic application of these
tabulated data. Rather, this character list is meant to demonstrate the phenetic
status of the single known A. garhi specimen with respect to features used to

features shared by A. aethiopicus, A. robustus, and A. boisei. Australopithecus garhi is

distinguished from A. africanus and other
early Homo species by its primitive frontal,
facial, palatal, and subnasal morphology.
Dental description. The postcanine dental size is remarkable, at or beyond the known
nonrobust and A. robustus extremes (Fig. 2).
The anterior dentition is also large, with I1
and canine breadths equivalent to or exceed-

evaluate early hominid fossils. Note that despite the large postcanine dentition,
no shared derived characters link A. garhi with A. robustus or A. boisei. The early
Homo column comprises specimens assigned by various authors to both H.
habilis and H. rudolfensis. Abbreviations are as follows: mod., moderate; asym.,
asymmetric; disp., disparate; sym., symmetric; rect., rectangular; para., parabolic;
var., variable; conv., convergent; div., divergent; ant., anterior; prom., prominent;
proc., procumbent; cont., continuous; interm., intermediate.
Males of

A.
afarensis

A. garhi
(n 5 1)

Canine to postcanine ratio

Incisor to postcanine ratio
Postcanine absolute size
UP3 occlusal outline

large
large
mod.
asym.

large
smaller
large
more
oval
absent
disp.
asym.
minor
thick

UP3 mesiobuccal line extension

Postcanine PM/M cusp wear
Canine lingual shape
Premolar molarization
Enamel thickness

frequent
disp.
asym.
none
moderately
thick

Ant. vertical thickness

Dental arcade shape
Posterior dental arcade
Ant. depth
UI2/UC diastema
Incisor alveoli relative to bicanine line

thin
rect.
conv.
shallow
common
ant.

thin
rect.
div.
shallow
present
ant.

C jugum
Ant. pillars
Inferolateral nasal aperture margin
UI2 root lateral to nasal aperture
Canine fossa
Maxillary fossula
Anterior zygomatic root position
Zygomaticoalveolar crest
Clivus contour
Subnasal prognathism
Incisor procumbency
Subnasal to intranasal contours
Separation of vomeral/ant. septal
insertion
Lateral ant. facial contour
Facial dishing

prom.
absent
sharp
lateral
present
absent
M1
arched
convex
strong
proc.
discrete
strong

prom.
absent
sharp
in line
present
absent
M1-P4
arched
convex
strong
proc.
discrete
strong

bipartite
absent

bipartite
absent

Frontal trigon
Costa supraorbitalis
Temporal lines frontal convergence
Postorbital constriction
Sagittal crest in male
Relative size of posterior temporalis
Parietal transverse expansion/tuber
Parietomastoid angle
Cranial capacity

present
present
mod.
mod.
present
large
absent
flared
small

present
present
mod.
mod.
present
interm.
absent
weak
small

Early Homo

A. africanus

A.
aethiopicus
(n 5 1)

A. robustus

A. boisei

Dentition
large
large
mod.
more oval

large
smaller
mod. to large
more oval

unknown
smaller?
large
oval

small
small
large
oval

small
small
large
oval

rare
disp.
asym.
minor
thick

rare
disp.
asym.
minor
thick

absent
flat
unknown
pronounced
hyperthick

absent
flat
more sym.
pronounced
hyperthick

absent
flat
more sym.
pronounced
hyperthick

thin
var.
div.
var.
absent
ant.

thick
rect.
conv.
shallow
absent
in line

thick
para.
div.
usually deep
absent
in line

thick
para.
div.
deep
absent
in line

prom.
present
var.
medial
present
absent
M1-P4
var.
flat
var.
var.
var.
weak

weak
absent
blunt
medial
absent
absent
P4
weak
concave
strong
proc.
cont.
weak

weak
present
var.
medial
absent
present
P4-P3
var.
flat to concave
weak
more vertical
cont.
weak

weak
absent
var.
medial
absent
absent
P4/M1-P3
var.
flat to concave
weak
more vertical
var.
weak

var.
absent

straight
dished

straight
dished

straight
dished

absent
interm.
weak
mod.
rare
interm.
absent
weak
small

present
present
strong
marked
present
large
absent
flared
small

present
present
strong
marked
present
small
absent
weak
slightly
enlarged

present
present
strong
marked
present
small
absent
weak
slightly
enlarged

Palate
thin
para.
div.
deep
rare
var.
Lower face
var.
absent
sharp
medial
var.
absent
M1-P4
arched
flat/convex
var.
var.
discrete
usually strong
var.
absent
Vault
absent
torus
weak
mod.
rare
interm.
present
weak
enlarged

www.sciencemag.org SCIENCE VOL 284 23 APRIL 1999

631

REPORTS
ing those of their largest known Australopithecus and early Homo homologs. Thus,
despite exceptional postcanine size, dental
proportions of the holotype deviate markedly
from the robust Australopithecus condition.
The canine-to-premolar/molar size ratios are
comparable to those of A. afarensis, A. africanus, and early Homo. Relative canine to
incisor alveolar length is most similar to that
of A. africanus. Postcanine wear, with developed angular facets and retention of buccal
cusp saliency, differs distinctively from the
robust Australopithecus pattern. The upper
P3 is more derived than that of A. afarensis
and most A. africanus specimens in exhibiting a reduced mesiobuccal crown quadrant
and a weak transverse crest. The buccolingual
narrowing of premolars and first molars often
seen in early Homo is absent.
Cranial description. The lower face is
prognathic, with procumbent incisors. Canine
roots are placed well lateral to the nasal
aperture margin. The premaxillary surface is
separated from the nasal floor by a blunt
ridge and is transversely and sagittally convex. The palate is vertically thin (;3 mm at
M1/M2 midline). The zygomatic roots originate above P4/M1. The dental arcade is Ushaped, with slightly divergent dental rows
(Fig. 3). The temporal lines encroach deeply
on the frontal, past the midsupraorbital position, and probably met anterior to bregma.
The postglabellar frontal squama is depressed
in a frontal trigon. The localized frontal sinus

is limited to the medial one-third of the supraorbital surface. Postorbital constriction is

marked. The parietal bones have a wellformed, bipartite, anteriorly positioned sagittal crest that divides above lambda. An endocast was made from the aligned parietals and
frontal and was completed by sculpting by R.
Holloway. Cranial capacity was about 450
cm3, as measured by water displacement.
Taxonomic discussion. There is no current
agreement about how many pre-erectus Homo
species should be recognized or even on how
the genus Homo should be defined. The traditional conservative definition emphasizes adaptive plateau. Ironically, by this definition, the
early Homo species H. rudolfensis and H. habilis might be better placed in Australopithecus,
as this would affiliate the major adaptive breakthroughs in anatomy and behavior that characterize H. erectus (ergaster) with the earliest
defined occurrence of Homo. If A. garhi proves
to be the exclusive ancestor of the Homo clade
(see discussion below), a cladistic classification
might assign it to genus Homo. Here we provisionally adopt the conservative, grade-sensitive
alternative, emphasizing its small brain and
large postcanine dentition by assigning the new
Bouri species to Australopithecus. This attribution as well as our diagnosis and description
may require emendation when additional individuals representing the species are recovered
and firm postcranial associations are established (6).
Although the Bouri Hata postcrania cannot

presently be assigned to the new species A.

garhi, they illuminate aspects of hominid evolution. The past few years have witnessed a rash
of attempts to estimate early hominid limb
length proportions from fragmentary and unassociated specimens. These specimens have
been used to generate a variety of functional
and phylogenetic scenarios. Accurate estimates
of the limb proportions of early hominids, however, must be confined to the very few specimens that actually preserve relevant elements,
such as the A.L. 288-1 (Lucy) specimen and
KNM-WT 15000. The new Bouri VP-12/1
specimen is only the third Plio-Pleistocene
hominid to provide reasonably accurate limb
length proportions. The Olduvai Hominid 62
specimen of Homo habilis has been erroneously
argued to show humerus-to-femur proportions
more primitive than those of Lucy (7, 8), but
its femur length cannot be accurately estimated.
Other studies of limb proportions in early Australopithecus species are based on unassociated
joints and not on actual (or even estimated)
limb lengths (7).
The postcranial remains recovered from
BOU-VP-11, -12, and -35 cannot be conclusively allocated to taxon. The BOU-VP-12/1
specimen features a humanlike humeral/femoral ratio (Fig. 4). This ratio may be an
important derivation relative to A. afarensis,
because it marks the earliest known appearance of the relative femoral elongation that
characterizes later hominids. However, as in
A. afarensis, the specimens brachial index is

Fig. 2. Dental size of A. garhi compared

with other early hominid taxa and specimens. (A) Canine breadth for various taxa.
(B) The square root of calculated (MD 3
BL) premolar area. (C) The square root of
calculated (MD 3 BL) second molar area.
(D) Canine breadth relative to postcanine
tooth size for various taxa. (E) Anterior
alveolar length (mesial I1 to distal C) relative to postcanine tooth size. In (A) through
(C), taxon means, standard deviations,
ranges, and sample sizes (in parentheses)
are given. All measures were taken by T.W.
and G.S. on originals except for A.L. 444-2
and A.L. 417-1 (A. afarensis) and A.L. 666-1
(Homo), which are from (2, 14). Dental metrics for the BOU-VP-12/130 specimen are as follows ( XX broken; parentheses 5 estimate; mesiodistal
measure reported first, followed by buccolingual): RI1 XX, (9.2); RI2 6.9, 6.8; RC 11.6, 12.9; RP3 (11.0), 16.0; RP4 XX, XX; RM1 XX, XX; RM2 (14.4), (17.7);
RM3 (15.2), 16.9; LI2 6.7, 7.0; LC 11.7, 12.9; LP3 (11.4), 16.0; LP4 (11.4), 16.0; LM1 (14.4), (16.5). M, A. afarensis; V, A. africanus; , Homo; }, A.
robustus; , A. boisei; v, BOU-VP-12/130.

632

23 APRIL 1999 VOL 284 SCIENCE www.sciencemag.org

REPORTS
apelike. This suggests that upper armto
lower arm ratios persisted into the basal
Pleistocene and that the first hominid with
modern forearm proportions was probably
Homo erectus (ergaster). Because the A. afarensis forearm was also long relative to both
the humerus and femur, the femur must have
elongated before forearm shortening in early
hominids.
The BOU-VP-11 ulna is from a larger
individual, as is the BOU-VP-35/1 humeral
shaft (estimated total humeral length 5 310
to 325 mm). The latter is absolutely longer
than the humerus of BOU-VP-12/1 but is less
rugose and probably bore a smaller deltopectoral crest. These differences (in both size and
rugosity) are well within the species ranges of
extant hominoids. If both humeri represent
the same taxon, they could reflect sexual
dimorphism, which would be comparable to
that currently seen in A. afarensis. However,
it is perilous to speculate on differences between only two specimens as they may reflect
only fluctuating intraspecific variation in
morphology and body mass.
The few and fragmentary nonrobust Turkana Basin hominids that span the 2.7 to 2.3 Ma
time range are similar to the Bouri specimens in
both size and aspects of morphology. Postca-

nine dental arcade length of BOU-VP-12/130 is

equivalent to that of Omo 75-14, whereas individual teeth of the smaller Middle Awash mandibles (GAM-VP-1/1 and BOU-VP-17/1) are
comparable in size to the smaller specimens of
the Omo nonrobust collection. It is also important that BOU-VP-17/1 exhibits a derived lower
P3 morphology (1) most similar to the Omo
nonrobust and early Homo conditions and a
dental arcade shape concordant with that of the
holotype of A. garhi.
On the basis of size, BOU-VP-12/130 is a
male. The craniodental size dimorphism documented for the closely related A. afarensis
and A. boisei therefore predicts smaller individuals in A. garhi. The biochronologically
contemporary and morphologically compatible BOU-VP-17/1 and GAM-VP-1/1 specimens are considerably smaller and may be
females. This would suggest a shift in either
or both body and dentognathic sizes to averages greater than in A. afarensis. More specimens are needed to test this hypothesis.
The discovery of A. garhi provides a strong
test of many phylogenetic hypotheses that have
addressed the relationships among Plio-Pleistocene hominid taxa. The South African species
A. africanus was once widely considered to be
the most primitive hominid. Discoveries of A.

afarensis at Hadar and Laetoli displaced A.

africanus. This more primitive sister species (A.
afarensis) was in turn supplanted when the
increasingly older and more primitive sister
taxa A. anamensis (9) and Ardipithecus ramidus
(10) were identified. However, the geometry of
post-afarensis hominid phylogeny continues to
be the focus of debate.
The position of A. africanus relative to the
emergence of the genus Homo has been particularly difficult to resolve, even in the face
of unduly elaborate phylogenetic analyses
(11). One reason for this difficulty is the
fundamental disagreement on whether early
Homo comprises one sexually dimorphic (H.

N
PA

L.

A.

8-1

28

WT
M- 00
KN 150

/1

12

U-

BO

MO

HO

cm

Fig. 3. The most complete palates of A. afarensis (A.L. 200-1a; canine reset) (A) and A. boisei
(OH-5) (B) compared with that of A. garhi (C and D). The photograph (David L. Brill 1999\Atlanta)
was mirror-imaged on midline. Australopithecus garhi has relatively large canines like A. afarensis
and absolutely large but morphologically nonrobust premolars and molars. Drawings L. Gudz.

Fig. 4. Probable stages in the progressive differentiation of hominid long bone proportions (all
bones shown to the same scale). The humerus
(top), antebrachium (middle), and femur (bottom) are of about equal length in chimpanzees
(Pan). Modern humans differ in two primary
ways. Although our humerus is virtually the same
length, the femur is elongated and the antebrachium is shortened. These changes appear to have
emerged fully by ;1.5 Ma in H. erectus [all three
limb segments are virtually complete in KNM-ER15000 (15)]. On the basis of the other two partial
skeletons in which long bone length can now be
reliably estimated, the modern human pattern
appears to have emerged in two stages: (i) elongation of the femur, which is intermediate in
length relative to the humerus in A.L. 288-1 but
exhibits modern proportions in BOU-VP-12/1;
and (ii) shortening of the antebrachium, which
retains primitive proportions in both specimens
(16). Drawings L. Gudz.

www.sciencemag.org SCIENCE VOL 284 23 APRIL 1999

633

REPORTS
Fig. 5. (A) A cladogram
s
us
us
us
us
us cus
depicting relationships
ec
ec
ec
cu hec
ec
e
us
e
h
h
h
h
h
c
t
t
t
t
t
t
e
i
th
among widely recogpi
pi
pi s
pi
pi
pi
ith s lo is op
lo s
lo
lo cu
lo
ip du tra ens tral sis alo
nized early hominid
ra nu
ra tus
ra pi
ra i
t
t
t
t
d
i
r
o 1.0
s io
s e
m
Ar am Aus am us ren ust hi Ausrica Ausbus
taxa, including the
Au eth
Au ois Ho
r
n A fa A ar
f
o
a
b
a
a
r
a
1.5
new species A. garhi.
g
Note that an addition2.0
al clade is required
when two contempo2.5
rary forms of early
Homo are recognized.
3.0
A variety of possible
3.5
cladograms have been
generated from the
4.0
data available in the
hominid fossil record,
4.5
but none of these satA
isfactorily resolve the
B
polychotomy illustrated here (11). This cladogram adds A. garhi
to the unresolved node.
(B) The chronological
relationships of early
hominid taxa. Age is
given in Ma. (C to F)
Alternative phylogenies
depicting possible relationships among early
C
D
E
F
hominid taxa. Note that
these alternatives do not exhaust the possibilities and that not all are entirely consistent with the cladogram. It is not presently possible to choose among these
alternatives.

habilis) or two (H. habilis and H. rudolfensis) species. Most phylogenetic efforts have
placed A. africanus as the link between A.
afarensis and early Homo. This hypothesis
has been widely, but not universally, accepted. Most predicted that a population of
A. africanus would be found in eastern
Africa when the 2.5 Ma gap there was filled
by fossil discoveries.
The 2.5 Ma A. garhi is derived toward megadontia from A. afarensis, but in cranial anatomy it is definitively not A. africanus. Neither is
it a representative of the contemporary A. aethiopicus. It is in the right place, at the right
time, to be the ancestor of early Homo, however
defined. Nothing about its morphology would
preclude it from occupying this position. The
close spatial and temporal association between
A. garhi and behaviors thought to characterize
later Homo provide additional circumstantial
support. The temporal and possible phylogenetic placements of various hominid taxa relative to the new species from Bouri are reviewed
in Fig. 5.
Plio-Pleistocene hominid phylogenetics is
bedeviled by atomization of functionally correlated character complexes that probably emanate from restricted genomic shifts as well as
inadequate fossil samples (particularly for early
Homo). Table 1 compiles characters available
for A. garhi and related taxa bearing on phylogenetic placement. The discovery of the KNMWT 17000 specimen of A. aethiopicus demonstrated the pervasiveness of homoplasy in hominid evolution (12). Specimens such as KNMER 1590, KNM-ER 1470, KNM-ER 1802,

634

Malawi UR 501, and Omo 75-14 make it obvious that some early Homo specimens exhibit
megadontia evolved in parallel with robust Australopithecus. Australopithecus garhi is certainly megadont, at least relative to craniofacial
size. However, its lack of derived robust characters leaves it as a sister taxon to Homo but
absent many derived Homo characters. A strictly cladistic analysis of available data has continually failed to resolve the issue of the position of A. africanus (11). The resulting currently unresolved polychotomy (Fig. 5) stems from
the fact that those characters most widely used
in early hominid phylogenetic systematics are
predominantly related to masticatory adaptation
and are known to be both interdependent and
susceptible to parallel evolution. Other characters such as cranial base flexion and craniofacial
hafting are even more poorly understood. The
atomization of such morphological complexes
has led to lengthy trait lists, but the valence of
the individual characters is clearly compromised. Such exercises have been useful in establishing the extensive homoplasy present
among early hominids, but such confirmation
only accentuates the precarious nature of phylogenetic reconstructions based on an incomplete and highly fragmentary fossil record.
Even a combination of all available temporal, spatial, and (circumstantial) behavioral evidence fails to resolve whether the origin of
Homo was from South African A. africanus or
east African A. afarensis (or both). We now
know that a nonrobust species derived from A.
afarensis persisted in eastern Africa until at
least 2.5 Ma. Only additional fossils will con-

firm whether this form participated in a rapid

evolutionary transition or transitions resulting
in an early form or forms of Homo. Such rapid
transition may be signaled by the recently described A.L. 266-1 palate from Hadar deposits
that are claimed to be 2.33 Ma (2). This palate
is more derived than that of A. garhi. If A. garhi
is the direct ancestor of early Homo, as represented by such younger specimens as KNM-ER
1590 and KNM-ER 1470, additional major
craniofacial changes must have occurred after
2.5 Ma, many of them as direct consequences
of brain enlargement. Novel behavioral shifts
associated with meat and marrow procurement
by means of lithic technology may have played
instrumental selective roles during this critical
and perhaps short period of evolution.
References and Notes

1. G. Suwa, T. D. White, F. C. Howell, Am. J. Phys.

Anthropol. 101, 247 (1996).
2. W. H. Kimbel et al., J. Hum. Evol. 31, 549 (1996); D. C.
Johanson and Y. Rak, Am. J. Phys. Anthropol. 103, 235
(1997).
3. A. Hill, S. Ward, A. Deino, G. Curtis, R. Drake, Nature
355, 719 (1992).
4. J. de Heinzelin et al., Science 284, 625 (1999).
5. B. A. Wood, Nature 355, 783 (1992).
6. The Chemeron temporal (3) has been called the
earliest Homo, but the only two characters cited in
support of this attribution (a medially positioned
mandibular fossa and a sharp petrous crest) are missing from the Bouri holotype, and neither provides
unambiguous evidence of brain expansion. Only additional discoveries will test whether Chemeron is a
Kenyan representative of A. garhi.
7. H. M. McHenry and L. R. Berger, J. Hum. Evol. 35, 1
(1998).
8. S. Hartwig-Scherer and R. D. Martin, ibid. 21, 439
(1991).

23 APRIL 1999 VOL 284 SCIENCE www.sciencemag.org

REPORTS
9. M. G. Leakey, C. S. Feibel, I. McDougall, A. C. Walker,
Nature 376, 565 (1995).
10. T. D. White, G. Suwa, B. Asfaw, ibid. 371, 306 (1994);
ibid. 375, 88 (1995).
11. A. T. Chamberlain and B. A. Wood, J. Hum. Evol. 16,
119 (1987); R. R. Skelton and H. M. McHenry, ibid. 23,
309 (1992); D. S. Strait, F. E. Grine, M. A. Moniz, ibid.
32, 17 (1997); R. R. Skelton and H. M. McHenry, ibid.
34, 109 (1997).
12. F. E. Grine, Ed., Evolutionary History of Robust Australopithecines (de Gruyter, New York, 1988); A. C.
Walker, R. E. Leakey, J. M. Harris, F. H. Brown, Nature
322, 517 (1986); H. M. McHenry, in Contemporary
Issues in Human Evolution, W. E. Meikle, F. C. Howell,
N. G. Jablonski, Eds. (California Academy of Sciences,
San Francisco, 1996), pp. 7792.
13. W. H. Kimbel, T. D. White, D. C. Johanson, Am. J. Phys.
Anthropol. 64, 337 (1984); G. Suwa et al., Nature
389, 489 (1997); P. V. Tobias, Olduvai Gorge, Volume
4: The Skulls, Endocasts and Teeth of Homo habilis
(Cambridge Univ. Press, Cambridge, 1991).
14. W. H. Kimbel, D. C. Johanson, Y. Rak, Nature 368, 449
(1994).
15. A. C. Walker and R. E. Leakey, Eds., The Nariokotome
Homo erectus Skeleton (Harvard Univ. Press, Cambridge, MA, 1993).
16. Femur and humerus length were virtually complete in
A.L. 288-1 [D. C. Johanson et al., Am. J. Phys. Anthropol. 57, 403 (1982)]. In BOU-VP-12/1, the femur
is preserved from the intersection of the medial
terminus of the neck with the (missing) femoral head
(proximally) to a point on the medial supracondylar
line just superior to the gastrocnemius impression
(distally). This distance was measured in a sex- and
species-balanced sample of Pan, Gorilla, and Homo
(N 5 60) and used to regress (least squares) femoral
length [correlation coefficient (r2) 5 0.952; 95%
confidence interval of estimate 5 60.28]. This regression computes the BOU-VP-12/1 femur at 348
mm. On anatomical grounds, we believe it to have
actually been slightly shorter (about 335 mm). Much
of the shaft of the BOU-12/1 humerus is preserved,
including the point of confluence between the diaphysis and the medial epicondylar apophysis and the
distalmost extent of the deltopectoral crest. This
distance was used to regress humeral length with the
same sample (length estimate 5 226 mm; r2 5
0.876; 95% confidence interval of estimate 5
60.40). On anatomical grounds, we estimate the
humerus to have been slightly longer (about 236
mm). Radial length was estimated for A.L. 288-1 with
multiple linear regressions from the same sample
(breadth distal articular surface; maximum diameter
radial head; length radial neck; r2 5 0.929; 95% confidence interval of estimate 5 60.29) and for BOU-VP12/1 (radial head to nutrient foramen; maximum diameter radial head; length radial neck; r2 5 0.937; 95%
confidence interval of estimate 5 60.27). These regressions estimate a length of 203 mm for A.L. 288-1 and
231 mm for BOU-VP-12/1. On anatomical grounds, the
BOU-VP-12/1 estimate appears correct. However, we
believe that the A.L. 288-1 radius is underestimated on
the basis of a lack of sufficient anatomical space with
which to accommodate all of the preserved pieces of
the bone. A regression limited to a sample of common
chimpanzees and bonobos (N 5 36) estimates a length
of 215 mm (r2 5 0.529; 95% confidence interval of
estimate 5 60.36). This result appears more probable.
Only exceptionally pronounced errors in any of the
above predictions would alter the conclusions made in
the legend of Fig. 3, nor are these conclusions altered by
regressions based only on single hominoid species.
17. The Middle Awash paleoanthropological project is multinational (13 countries), interdisciplinary research codirected by B.A., Y. Beyene, J. D. Clark, T.W., and G.
WoldeGabriel. The research reported here was supported by the NSF. We thank N. Tahiro and A. Abdo for their
assistance in naming the new species. We thank Y.
Haile-Selassie for discovery of the BOU-VP-12/130 holotype and H. Gilbert and D. DeGusta for field and
illustrations work. R. Holloway kindly allowed us to cite
his BOU-VP-12/130 cranial capacity estimate. L. Gudz
made the palate and postcranial drawings. D. Brill made
the photographs. P. Reno provided comparative primate

data. The Japan Ministry of Education, Science, Sports

and Culture provided support to G.S. We thank the
Ethiopian Ministry of Information and Culture, the Centre for Research and Conservation of the Cultural Heritage, and the National Museum of Ethiopia. We thank
the Afar Regional Government and the Afar people of

the Middle Awash for permission and support. We

thank the many individuals who contributed to the
camp, transport, survey, excavation, and laboratory
work that stands behind the results presented.
25 February 1999; accepted 30 March 1999

Electron Solvation in Finite

Systems: Femtosecond
Dynamics of Iodidez( Water)n
Anion Clusters
L. Lehr,* M. T. Zanni,* C. Frischkorn, R. Weinkauf,
D. M. Neumark
Electron solvation dynamics in photoexcited anion clusters of I2(D2O)n54 6
and I2(H2O)4 6 were probed by using femtosecond photoelectron spectroscopy (FPES). An ultrafast pump pulse excited the anion to the cluster analog of
the charge-transfer-to-solvent state seen for I2 in aqueous solution. Evolution
of this state was monitored by time-resolved photoelectron spectroscopy using
an ultrafast probe pulse. The excited n 5 4 clusters showed simple population
decay, but in the n 5 5 and 6 clusters the solvent molecules rearranged to
stabilize and localize the excess electron, showing characteristics associated
with electron solvation dynamics in bulk water. Comparison of the FPES of
I2(D2O)n with I2(H2O)n indicates more rapid solvation in the H2O clusters.
A free electron can be trapped by solvent reorientation in polar solvents such as ammonia (1)
or water (2). These solvated electrons play an
important role in condensed phase chemistry,
including radiation chemistry, electron transfer,
and charge-induced reactivity. A microscopic
understanding of the electron-solvent and solvent-solvent interactions that govern electron
solvation is therefore a fundamental and challenging problem. These considerations have
motivated femtosecond time-resolved studies
that have demonstrated rich and complex dynamics after electronic excitation of electrons in
water (3, 4).
To gain a complementary perspective on
this problem, we studied solvated electron
dynamics in finite clusters and compared
these results with our understanding of bulk
solvation phenomena. We used two-photon
anion femtosecond (10215 s) photoelectron
spectroscopy (FPES) (5, 6) to study electron
solvation dynamics in the mass-selected anion clusters I2(D2O)n and I2(H2O)n in order
to address the following questions: (i) What
Department of Chemistry, University of California,
Berkeley, CA 94720, USA and Chemical Sciences Division, Lawrence Berkeley National Laboratory, Berkeley, CA 94720, USA.
*These authors contributed equally to this work.
Permanent address: Institut fur Physikalische und Theoretische Chemie, Technischen Universitat Munchen,
D-85748 Garching, Germany.
To whom correspondence should be addressed. Email: dan@radon.cchem.berkeley.edu

is a minimum solvent cluster size needed to

solvate an electron? (ii) What is a typical time
scale for solvent reorientation in a cluster?
(iii) What type of solvent motion is involved
in electron solvation dynamics?
Aqueous solutions of I2 exhibit broad
electronic bands in the ultraviolet (UV) corresponding to electron ejection from I2 into
the solvent (7), known as charge-transferto-solvent (CTTS) states. Excitation of these
states is an elegant means of generating solvated electrons, as was first demonstrated by
Jortner (8). The dynamics of these states have
been investigated by Eisenthal (9), Gaudeul
(10), Bradforth (11), and their co-workers, all
of whom excited the CTTS states to inject an
electron into the water and then followed the
subsequent electron solvation dynamics by
femtosecond absorption spectroscopy. These
experimental studies along with simulations
by Sheu and Rossky (12) and Staib and Borgis (13) show that excitation of the lowest
energy CTTS band results in the generation
of fully solvated electrons on a 200-fs time
scale (11). Once generated, these electrons
thermalize with the solvent molecules, and
some are then lost through geminate recombination with the neutral halogen atom over a
time scale of tens of picoseconds.
The issue of how the CTTS bands manifest themselves in finite clusters was first
addressed in experiments by Johnson and
co-workers (14), in which a diffuse absorption band was seen just above the detachment

www.sciencemag.org SCIENCE VOL 284 23 APRIL 1999

635